Partial migration is common in a large variety of taxa in seasonally variable environments. Understanding the mechanisms underlying migration is important, as migration affects individual fi tness. Migratory herbivores bene fi t from delayed forage maturation and hence higher food quality during migra- tion and at their summer range, termed the forage maturation hypothesis (FMH). The link between diet quality and rumination time allows migrants eating a higher quality diet to spend less time on rumination, and they can thus allocate more time to additional feeding. However, such an argument implicitly assumes that deer are energy maximizers, while studies have reported also time minimization strategies under risk of predation. Male and female distributions are limited by different factors linked to both body size differ- ences and reproductive strategies, but there is no study investigating differences in activity pattern accord- ing to the individual migratory patterns for male and female deer. We here unify the FMH with the hypotheses predicting sex-speci fi c time allocation strategies. To test predictions of sex-speci fi c activity of resident and migratory red deer ( Cervus elaphus ), we analyzed activity data of 286 individuals that were fi tted with GPS collars from a population in western Norway. While migrants were more active during the migration itself, we found no differences in activity pattern between migrant and resident deer during the main growth season, neither in terms of proportion of daily time active nor in terms of daily mean movement speed, thus rejecting that deer were energy maximizers. Overall, we found that females were more active during the main growth season even after controlling for body size differences. These patterns are consistent with patterns predicted from sexual segregation theory linked to the reproductive strategy hypothesis. Our study highlights how the understanding of migration can be advanced by considering it in the context of different reproductive strategies of males and females.

1. Population-level management is difficult to achieve if wildlife routinely crosses administrative boundaries, as is particularly frequent for migratory populations. However, the degree of mismatch between management units and scales at which ecological processes operate has rarely been quantified. Such insight is vital for delimiting functional population units of partially migratory species common in northern forest ecosystems. 2. We combined an extensive dataset of 412 GPS-marked red deer (Cervus elaphus) across Norway with information on the size and borders of two administrative levels, the governmental level (municipality) and landowner level (local management units, LMUs), to determine the timing and scale of mismatch between animal space use and management units. We analysed how landscape characteristics affected the use of management units and the timing and likelihood of crossing borders between them, in an effort to delineate more appropriate units in various landscapes. 3. Median municipality size could potentially cover 70% of female and 62% of male annual ranges, while only 12% and 4% of LMUs were expansive enough to accommodate migratory routes in females and males, respectively. Red deer migrate along elevational gradients and are more likely to find both suitable lowland winter habitat and higher summer habitat within management units with variable topography. Consistent with this, the likelihood of border crossing decreased with increasing diversity of elevations. 4. Synthesis and applications. We demonstrate a considerable mismatch between animal space use and management units. Far-ranging movements and frequent administrative border crossings during autumn migration coincides with the period of active management (hunting season). Our study also highlights that, due to extensive movements of males, coordination of management aims may provide a more realistic avenue than increasing sizes of local management units. A more general insight is that the degree of mismatch between range use and management units depends on the season and landscape type. This needs to be accounted for when delimitating functional population units of migratory populations.

Large herbivores gain nutritional benefits from following the sequential flush of newly emergent, high- quality forage along environmental gradients in the landscape, termed green wave surfing. Which landscape characteristics underlie the environmental gradi-ent causing the green wave and to what extent landscape characteristics alone explain individual variation in nutritional benefits remain unresolved questions. Here, we com-bine GPS data from 346 red deer (Cervus elaphus) from four partially migratory popula-tions in Norway with the satellite- derived normalized difference vegetation index (NDVI), an index of plant phenology. We quantify whether migratory deer had access to higher quality forage than resident deer, how landscape characteristics within sum-mer home ranges affected nutritional benefits, and whether differences in landscape characteristics could explain differences in nutritional gain between migratory and resident deer. We found that migratory red deer gained access to higher quality forage than resident deer but that this difference persisted even after controlling for land-scape characteristics within the summer home ranges. There was a positive effect of elevation on access to high- quality forage, but only for migratory deer. We discuss how the landscape an ungulate inhabits may determine its responses to plant phenol-ogy and also highlight how individual behavior may influence nutritional gain beyond the effect of landscape.

As global warming advances, there is a growing concern about the impact of extreme weather events on ecosystems. In the Arctic, more frequent unseasonal warm spells and rain-on- snow events in winter cause changes in snow-pack properties, including ground icing. Such extreme weather events are known to have severe effects across trophic levels, for instance, causing die-offs of large herbivores. However, the extent to which individuals and populations are able to buffer such events through behavioral plasticity is poorly understood. Here, we analyze responses in space use to rain-on- snow and icing events, and their fitness correlates, in wild reindeer in high-Arctic Svalbard. Range displacement among GPS-collared females occurred mainly in icy winters to areas with less ice, lower over-winter body mass loss, lower mortality rate, and higher subsequent fecundity, than the departure area. Our study provides rare empirical evidence that mammals may buffer negative effects of climate change and extreme weather events by adjusting behavior in highly stochastic environments. Under global warming, behavioral buffering may be important for the long-term population persistence in mobile species with long generation time and therefore limited ability for rapid evolutionary adaptation. Arctic; climate; GPS; ice; ideal-free distribution; migration; movement; Rangifer tarandus platyrhynchus; space use; Svalbard; Svalbard reindeer; time-to-event analysis.